Seasonal and annual differences in the foraging ecology of two gull species breeding in sympatry and their use of fishery discards

Niche segregation between similar species will result from an avoidance of competition but also from environmental variability, including nowadays anthropogenic activities. Gulls are among the seabirds with greater behavioural plasticity, being highly opportunistic and feeding on a wide range of prey, mostly from anthropogenic origin. Here, we analysed blood and feather stable isotopes combined with pellet analysis to investigate niche partitioning between Audouin's gull Larus audouinii and yellow‐legged gull Larus michahellis breeding in sympatry at Deserta Island, southern Portugal, during 2014 and 2015. During the breeding season there was considerable overlap in the adults’ diet, as their stable isotope values of blood and primary feather (P1) did not differ, and their pellets were comprised mainly by marine fish species. However, Audouin's gulls presented higher occurrences of pelagic fish, while yellow‐legged gulls fed more on demersal fish, insects, and refuse. SIAR mixing models also estimated a higher proportion of demersal fish in the diet of yellow‐legged gulls. We also found differences between the two gull species in chicks’ feathers, suggesting that Audouin's gull adults selected prey with lower carbon isotope values to feed their young. Secondary feather (S8) of Audouin's gull presented higher isotope values compared to yellow‐legged gulls, indicating different foraging areas (δ¹³C) and/ or trophic levels (δ¹⁵N) between the two species in the non‐breeding season. During both the all‐year and non‐breeding periods the yellow‐legged gull showed a broader isotopic niche width than Audouin's gull in 2013, and in 2014 the two gull species exhibited different isotopic niche spaces. Our study suggests that both gull species foraged in association with fisheries during the breeding season. In this sense, a discard ban implemented under the new European Union Common Fisheries Policy may lead to a food shortage, therefore future research should closely monitor the population dynamics of Audouin's and yellow‐legged gulls.     

Effects of ethanol storage and lipid and urea extraction on δ15N and δ13C isotope ratios in a benthic elasmobranch,the bluespotted maskray Neotrygon kuhlii

Ethanol storage and lipid and urea extraction had no effect on bluespotted maskray Neotrygon kuhlii muscle δ¹³C values whereas urea‐removal and ethanol storage increased δ¹⁵N values. Results presented here show a significant δ¹⁵N increase post‐urea removal and provide additional support for this approach in future elasmobranch stable‐isotope analysis (SIA) studies. Further experimental work on other elasmobranch species is needed to assess extraction and preservation effects on stable‐isotope (SI) values.     

The influence of lipid‐extraction and long‐term DMSO preservation on carbon (δ13C) and nitrogen (δ15N) isotope values in cetacean skin

Stable isotope analysis (SIA) has rapidly become a useful tool to study the ecology of wild animal populations, especially for elusive, wide‐ranging predators like marine mammals. The development of projectile biopsy techniques resulted in the collection of thousands of cetacean tissue samples that were archived in a dimethyl sulfoxide (DMSO) solution for long‐term, multidecadal preservation. Here we examine the influence of DMSO preservation on carbon (δ¹³C) and nitrogen (δ¹⁵N) values by comparing a set of paired delphinid skin samples stored frozen without preservative and in DMSO for up to 22 yr. Treatment of paired frozen and DMSO‐preserved skin in a 2:1 chloroform:methanol solution yielded similar δ¹³C and δ¹⁵N values, revealing that DMSO and lipid contamination have similar isotopic effects on skin, and that these effects can be removed using routine lipid‐extraction methods. Further, amino acid concentrations in DMSO‐preserved and frozen skin tissue were similar, providing independent evidence of minimal protein alteration due to preservation. Access to a rich archive of skin samples preserved in DMSO will expand our ability to examine temporal and spatial variability in the isotope values of cetaceans, which will aid our understanding of how their ecology has been influenced by historical changes in environmental conditions.     

The effects of tissue type and body size on δ13C and δ15N values in parrotfish (Labridae) from Zanzibar,Tanzania

Differences between the stable isotopic ratios (δ¹³C and δ¹⁵N) of two tissues (blood and muscle) from four species of East African coral reef parrotfishes (family: Labridae, tribe: Scarini) were analysed across a broad spectrum of body sizes. Comparison of isotopic ratios between the tissues allowed the assessment of using blood as an alternative tissue to muscle. In 2010–2011, constant differences between tissues (δ_blood minus δ_muscle) were found across a broad range of sampled fish lengths. Linear relationships between the tissues, specific for an isotope, indicate that constants could be generated for converting blood isotope into muscle isotope values. Only one species, Chlorurus sordidus, displayed an inconsistent difference between tissues in δ¹⁵N, indicating that this ratio was dependent on fish length. The δ¹³C of both tissues was positively related linearly to fish length for three species, while δ¹⁵N showed no relationship with body length. The results are interpreted as indicating dietary consistency over days to weeks, the time of tissue turnover for blood and muscle, respectively. Lastly, differences among the species, even closely related species, show that the generation of tissue conversion constants is species‐specific.     

Comparative response of δ13C, δ18O and δ15N in durum wheat exposed to salinity at the vegetative and reproductive stages

This study compared the performance of the stable isotope composition of carbon (δ¹³C), oxygen (δ¹⁸O) and nitrogen (δ¹⁵N) by tracking plant response and genotypic variability of durum wheat to different salinity conditions. To that end, δ¹³C, δ¹⁸O and δ¹⁵N were analysed in dry matter (dm) and the water‐soluble fraction (wsf) of leaves from plants exposed to salinity, either soon after plant emergence or at anthesis. The δ¹³C and δ¹⁸O of the wsf recorded the recent growing conditions, including changes in evaporative conditions. Regardless of the plant part (dm or wsf), δ¹³C and δ¹⁸O increased and δ¹⁵N decreased in response to stress. When the stress conditions were established just after emergence, δ¹⁵N and δ¹³C correlated positively with genotypic differences in biomass, whereas δ¹⁸O correlated negatively in the most severe treatment. When the stress conditions were imposed at anthesis, relationships between the three isotope signatures and biomass were only significant and positive within the most severe treatments. The results show that nitrogen metabolism, together with stomatal limitation, is involved in the genotypic response to salinity, with the relative importance of each factor depending on the severity and duration of the stress as well as the phenological stage that the stress occurs.     

Effects of decomposition and storage conditions on the δ13C and δ15N isotope values of killer whale (Orcinus orca) skin and blubber tissues

Several different factors in the collection and preservation of whale skin and blubber samples were examined to determine their effect on the results obtained by stable nitrogen and carbon isotope (δ¹⁵N and δ¹³C) analysis. Samples of wet killer whale skin retained their original stable isotope values for up to 14 d at 4°C or lower. However, decomposition significantly changed the δ¹⁵N value within 3 d at 20°C. Storage at ?20°C was as effective as ?80°C for the preservation of skin and blubber samples for stable isotope analysis for at least a year. By contrast, once a skin sample had been freeze‐dried and lipid extracted, the stable isotope values did not change significantly when it was stored dry at room temperature for at least 12 mo. Preservation of whale skin samples for a month in DMSO‐salt solution, frozen or at room temperature, did not significantly change the δ¹⁵N and δ¹³C values of lipid extracted tissues, although the slight changes seen could influence results of a study if only small changes are expected.     

Relationships between growth traits and scale stable isotopes (δ13C, δ15N) of adult chum salmon Oncorhynchus keta in Hokkaido,Japan

Stable isotope analysis (SIA) in combination with growth analysis using scales collected from adult chum salmon Oncorhynchus keta migrating back to Hokkaido, Japan, was performed to describe the variation of isotopic composition of carbon (δ¹³C) and nitrogen (δ¹⁵N) in scales and to examine relationships with growth traits [age, fork length (FL), and relative growth ratio in the last growth period [(RGR^last)]. Scale stable isotope (SI) values in 3‐ to 6‐year‐old fish were highly variable, ranging from ?17.6‰ to ?14.3‰ for δ¹³C and 9.5‰ to 13.4‰ for δ¹⁵N. The δ¹⁵N was positively correlated with FL, and this tendency may indicate changes in trophic level with growth. Significant effect was not detected between δ¹⁵N and RGR^last, it can be inferred that factors potentially yielding high δ¹⁵N may not necessarily result in higher growth rates. No trend found between FL and δ¹³C may imply that there is no clear segregation in feeding locations between the 3‐ to 6‐year groups. This study provided basic information for scale SI values of chum salmon adults and indicated that SIA using scales could be a new approach to elucidating the trophic ecology of chum salmon.     

A stable isotopic investigation into the causes of decline in a sub-Antarctic predator, the rockhopper penguin Eudyptes chrysocome

The rockhopper penguin (Eudyptes chrysocome) is a conspicuous apex marine predator that has experienced marked population declines throughout most of its circumpolar breeding distribution. The cause(s) for the declines remain elusive, but the relatively large spatio‐temporal scale over which population decreases have occurred implies that ecosystem‐scale, at‐sea factors are likely to be involved. We employ stable isotope analyses of carbon (¹³C/¹²C, expressed as δ¹³C) and nitrogen (¹⁵N/¹⁴N, δ¹⁵N) in time‐series of rockhopper penguin feather samples, dating back to 1861, in order to reconstruct the species' ecological history. Specifically, we examine whether rockhopper penguin population decline has been associated with a shift towards lower primary productivity in the ecosystem in which they feed, or with a shift to a diet of lower trophic status and lower quality, and we use long‐term temperature records to evaluate whether shifts in isotope ratios are associated with annual variations in sea surface temperature. Having controlled temporally for the Suess Effect and for increases in CO₂ concentrations in seawater, we found that overall, δ¹³C signatures decreased significantly over time in rockhopper penguins from seven breeding sites, supporting the hypothesis that decreases in primary productivity, and hence, carrying capacity, for which δ¹³C signature is a proxy, have been associated with the decline of penguin populations. There was some evidence of a long‐term decline in δ¹⁵N at some sites, and strong evidence that δ¹⁵N signatures were negatively related to sea surface temperatures across sites, indicative of a shift in diet to prey of lower trophic status over time and in warm years. However, a site‐by‐site analysis revealed divergent isotopic trends among sites: five of seven sites exhibited significant temporal or temperature‐related trends in isotope signatures. This study highlights the utility of stable isotope analyses when applied over relatively long timescales to apex predators.     

The effects of Paraloid B‐72 and Butvar B‐98 treatment and organic solvent removal on δ13C, δ15N,and δ18O values of collagen and hydroxyapatite in a modern bone

Stable isotopes in bones are a powerful tool for diet, provenance, climate, and physiological reconstructions, but necessarily require well‐preserved specimens unaltered by postmortem diagenesis or conservation practices. This study examines the effects of Paraloid B‐72 and Butvar B‐98, two common consolidants used in field and museum conservation, on δ¹³C, δ¹⁵N, and δ¹⁸O values from bone collagen and hydroxyapatite. The effects of solvent removal (100% acetone, 100% ethanol, 9:1 acetone:xylenes, 9:1 ethanol:xylenes) and drying methods (ambient air, vacuum, oven drying at 80°C) were also examined to determine if bones treated with these consolidants can successfully be cleaned and used for stable isotope analyses. Results show that introduction of Paraloid B‐72 or Butvar B‐98 in 100% acetone or 100% ethanol, respectively, with subsequent removal by the same solvents and drying at 80°C facilitates the most successful removal of consolidants and solvents. The δ¹³C values in collagen, δ¹⁵N in collagen, δ¹⁸O in hydroxyapatite phosphate, and δ¹³C in hydroxyapatite structural carbonate were unaltered by treatments with Paraloid or Butvar and subsequent solvent removal. The δ¹⁸O in hydroxyapatite structural carbonate showed nonsystematic variability when bones were treated with Paraloid and Butvar, which is hypothesized to be a result of hydroxyl exchange when bones are exposed to consolidants in solution. It is therefore recommended that δ¹⁸O in hydroxyapatite structural carbonate should not be used in stable isotope studies if bones have been treated with Paraloid or Butvar. Am J Phys Anthropol 157:330–338, 2015. © 2015 Wiley Periodicals, Inc.     

Variability in stable isotopes of snowy owl feathers and contribution of marine resources to their winter diet

The snowy owl is an elusive arctic predator known for its nomadic behaviour. Satellite tracking has revealed that some adult snowy owls could make an extensive use of the marine environment during the non‐breeding season. However, the relative contribution of marine resources to their diet is unknown. Stable isotope analyses can be useful to document the diet of mobile animals during periods of the year when individuals are less accessible. This study aimed to assess variation in isotopic values (δ¹³C and δ¹⁵N) of various feather types, and the usefulness of feathers to determine the contribution of the marine environment to the winter diet of snowy owls captured in summer. We sampled feathers coming from 6 body regions of 18 breeding females at two sites in the eastern Canadian Arctic in 2013 and 2014. Prior to analyses, diet‐tissue discrimination factors of snowy owl feathers were established in captivity. Variability in isotopic values among feather types was relatively low and pairwise correlations in isotopic values between feathers on the same individual were variable and often low, which suggests differences in the diet at the time when various feathers were synthesized. Diet reconstruction models detected a contribution of marine sources to snowy owl feathers ranging from 4 to 19% among feather types. However, the marine contribution was highly variable when single feathers were examined within individuals, ranging from 3 to 71%. This indicated that no single feather type could be used alone to reliably infer the contribution of marine resources to the winter diet of owls, possibly due to a high variability in the timing and sequence of molt. For asynchronous molters like snowy owls, we recommend sampling multiple feathers from various body regions, excluding wing feathers, to investigate winter diet or habitat use.     

Migratory connectivity in a declining bird species: using feather isotopes to inform demographic modelling

Aim Conservation programmes for endangered migratory species or populations require locating and evaluating breeding, stopover and wintering areas. We used multiple stable isotopes in two endangered European populations of wrynecks, Jynx torquilla L., to locate wintering regions and assess the degree of migratory connectivity between breeding and wintering populations. Location Switzerland and Germany. Methods We analysed stable nitrogen (δ¹⁵N), carbon (δ¹³C) and hydrogen (δD) isotopes from wing feathers from two populations of wrynecks to infer their wintering origins and to assess the strength of migratory connectivity. We tested whether variation in feather isotopic values within the Swiss population was affected by bird age and collection year and then considered differences in isotopic values between the two breeding populations. We used isotopic values of summer‐ and winter‐grown feathers to estimate seasonal distributions. Finally, we calculated a species‐specific δD discrimination factor between feathers and mean annual δD values to assign winter‐grown feathers to origin. Results Bird age and collection year caused substantial isotopic variation in winter‐grown feathers, which may be because of annually variable weather conditions, movements of birds among wintering sites and/or reflect asynchronous moulting or selection pressure. The large isotopic variance in winter‐grown feathers nevertheless suggested low migratory connectivity for each breeding population, with partially overlapping wintering regions for the two populations. Main conclusions Isotopic variance in winter‐grown feathers of two breeding populations of wrynecks and their geographical assignment point to defined, albeit overlapping, wintering areas, suggesting both leapfrog migration and low migratory connectivity. On this basis, integrative demographic models can be built looking at seasonal survival patterns with links to local environmental conditions on both breeding and wintering grounds, which may elucidate causes of declines in migratory bird species.     

Carbon (δ13C) and nitrogen (δ15N) stable isotope composition in plant and soil in Southern Patagonia's native forests

Stable isotope natural abundance measurements integrate across several biogeochemical processes in ecosystem N and C dynamics. Here, we report trends in natural isotope abundance (δ¹³C and δ¹⁵N in plant and soil) along a climosequence of 33 Nothofagus forest stands located within Patagonia, Southern Argentina. We measured 28 different abiotic variables (both climatic variables and soil properties) to characterize environmental conditions at each of the 33 sites. Foliar δ¹³C values ranged from ?35.4‰ to ?27.7‰, and correlated positively with foliar δ¹⁵N values, ranging from ?3.7‰ to 5.2‰. Soil δ¹³C and δ¹⁵N values reflected the isotopic trends of the foliar tissues and ranged from ?29.8‰ to ?25.3‰, and ?4.8‰ to 6.4‰, respectively, with no significant differences between Nothofagus species (Nothofagus pumilio, Nothofagus antarctica, Nothofagus betuloides). Principal component analysis and multiple regressions suggested that mainly water availability variables (mean annual precipitation), but not soil properties, explained between 42% and 79% of the variations in foliar and soil δ¹³C and δ¹⁵N natural abundance, which declined with increased moisture supply. We conclude that a decline in water use efficiency at wetter sites promotes both the depletion of heavy C and N isotopes in soil and plant biomass. Soil δ¹³C values were higher than those of the plant tissues and this difference increased as annual precipitation increased. No such differences were apparent when δ¹⁵N values in soil and plant were compared, which indicates that climatic differences contributed more to the overall C balance than to the overall N balance in these forest ecosystems.     

Tracing site‐specific isotopic signatures along a Blue Tit Cyanistes caeruleus food chain

Food chains culminating with temperate insectivorous passerines are well described, yet whether trophic webs can be site‐specific remains a largely unexplored question. In the case of site‐ or habitat‐specificity of food webs, stable isotope signatures of bird feathers may enable assignment of unmarked individuals to a site or a habitat of origin. We address this question in landscapes that include contrasting forest habitat patches with either deciduous Downy Oak Quercus humilis or evergreen Holm Oak Quercus ilex as dominant tree species. First, we examine the spatial variation across habitats and sites in the stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) along the oak leaf–Tortrix moth Tortrix viridana caterpillar–Blue Tit Cyanistes caeruleus food chain. Secondly, we assess whether the isotopic signatures allow for correct assignment of individual birds to their site or habitat of origin. At the scale of the landscape, stable isotope values enabled identification of the different components of the Blue Tit food chain: from oak leaves to Blue Tit nestlings and yearling birds. However, isotopic signatures were site‐specific (i.e. geographical) more than habitat‐specific (i.e. deciduous vs. evergreen oaks). Discriminant analyses correctly assigned 85% of nestlings and 83% of resident yearling birds, indicating a pronounced effect of site on Blue Tit feather isotopic signatures. We thereby demonstrate that isotopes reflect a stronger association of locally born birds to the local features of their habitat than that of un‐ringed yearling birds, whose plumage may have grown while in a wider geographical area. This study provides evidence of site‐specific isotopic signatures from oak leaves to Blue Tit feathers at a fine spatial scale.     

On the use of stable oxygen isotope (δ18O) measurements for tracking avian movements in North America

Tracking migratory animals has benefitted using measurements of naturally occurring stable isotopes of hydrogen (δ²H) in keratinous tissues such as hair and feathers to link animal origins to continental patterns or isoscapes of δ²H in precipitation. However, for most taxa, much less information exists on the use of stable oxygen isotope ratios (δ¹⁸O) despite the fact that δ²H and δ¹⁸O are strongly linked in environmental waters through the meteoric relationship and the possibility of using both isotopes to infer greater information on origins and climatic conditions where tissues are grown. A fundamental requirement of using stable isotopes to assign individuals and populations to origins is the development of a rescaling function linking environmental food web signals to the tissue of interest and for birds, this has not been carried out. Here, we derived the relationship between H and O isotopes in known source feathers of 104 individuals representing 11 species of insectivorous passerines sampled across the strong precipitation isoscape of North America. We determined again a strong expected relationship between feather δ²H (δ²H_f) and long‐term amount‐weighted precipitation δ²H (δ²H_p; r² = 0.77), but the corresponding relationship between δ¹⁸O_f and δ¹⁸O_p was poor (r² = 0.32) for the same samples. This suggests that δ²H measurements are currently more useful for assignment of insectivorous songbirds to precipitation isoscapes but does not preclude other uses of the δ¹⁸O_f data. Currently, mechanisms responsible for the decoupling of H and O isotopes in food webs is poorly known, and we advocate a much broader sampling of both isotopes in the same keratinous tissues across precipitation isotope gradients and across taxa to resolve this issue and to increase the power of using water isotopes to track migratory animals.     

Nonlethal sampling for stable isotope analysis of juvenile Chinese sturgeon (Acipenser sinensis): Comparing δ13C and δ15N signatures in muscle and fin tissues

We compared δ¹³C and δ¹⁵N values of muscle with fin from juvenile Chinese sturgeon (Acipenser sinensis), to evaluate the feasibility of using nonlethal (fin) as an alternative to lethal (muscle) sampling. Size and lipid effect on the relationship between fin and muscle were also investigated. Dorsal muscle (DM) and fin clip (FC) were collected from A. sinensis with different body length (120–373 mm) in the Yangtze Estuary for isotope analysis. The result showed that (1) muscle isotope values could estimated by the values of fin, from either use the regression model (δ¹³C_DM = 0.939 × FC ? 2.577; δ¹⁵N_DM = 0.737 × FC + 4.638) or constants factors (δ¹³C_DM = δ¹³C_FC ? 1.27; δ¹⁵N_DM = δ¹⁵N_FC + 0.59); (2) no size‐based relationships with δ¹³C and δ¹⁵N from either fin or muscle; (3) lipid extraction significantly improving the fin and muscle regression model fit for both δ¹³C and δ¹⁵N values. Therefore, this study support the use of nonlethal fin tissues for isotope analysis of juvenile A. sinensis, and will allow trophic studies to avoid the effect of lipid accumulation from muscle.     

Environmental and species‐specific controls on δ13C and δ15N in dominant woody plants from central‐western Argentinian drylands

Spatial variation in mean annual precipitation is the principal driver of plant water and nitrogen status in drylands. The natural abundance of carbon stable isotopes (δ¹³C) in photosynthetic tissues of C3 plants is an indicator of time‐integrated behaviour of stomatal conductance; while that of nitrogen stable isotopes (δ¹⁵N) is an indicator of the main source of plant N (soil N vs. atmospheric N₂). Previous studies in drylands have documented that plant δ¹³C and δ¹⁵N values increase with decreasing mean annual precipitation due to reductions in stomatal conductance, and soil enriched in ¹⁵N, respectively. However, evidence for this comes from studies focused on stable isotopes measurements integrated at the plant community level or on dominant plants at the site level, but little effort has been made to study C and N isotope variations within a species growing along rainfall gradients. We analysed plant δ¹³C, δ¹⁵N and C/N values of three woody species having different phenological leaf traits (deciduous, perennial and aphyllous) along a regional mean annual precipitation gradient from the central‐western Argentinian drylands. Noticeably, plant δ¹³C and δ¹⁵N values in the three woody species did not increase towards sites with low precipitation or at the start of the growing season (drier period), as we expected. These results suggest that environmental factors other than mean annual precipitation may be affecting plant δ¹³C and δ¹⁵N. The short‐term environmental conditions may interact with species‐specific plant traits related to water and nitrogen use strategies and override the predictive influence of the mean annual precipitation on plant δ¹³C and δ¹⁵N widely reported in drylands.     

Identifying critical habitat of the endangered vaquita (Phocoena sinus) with regional δ13C and δ15N isoscapes of the Upper Gulf of California,Mexico

The vaquita (Phocoena sinus) is the world's most endangered cetacean and has experienced a 60% reduction in the size of its population in the past decade. Knowledge of its basic ecology is essential for developing successful management plans to protect and conserve this species. In this study, we identified vaquita foraging areas by creating an isoscape of the Upper Gulf of California (UGC) based on sediment and zooplankton carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values. Our results confirm that this species is confined to the western region of the UGC, which is characterized by relatively high δ¹⁵N values (sediments: 10.2‰ ± 2.0‰, zooplankton: 15.8‰ ± 1.3‰), higher sea surface temperatures (～16°C–25°C), higher concentrations of silt in sediments, and the highest turbidity. In contrast, the eastern region of the UGC had relatively low sediment (7.7‰ ± 2.4‰) and zooplankton (14.6‰ ± 1.0‰) δ¹⁵N values, and the highest concentrations of sand in sediments. Our approach is an effective use of marine isoscapes over a small spatial scale (<200 km) to identify the environmental characteristics that define the critical habitat for an extremely endangered marine mammal.     

Metabolic origin of δ15N values in nitrogenous compounds from Brassica napus L. leaves

Nitrogen isotope composition (δ¹⁵N) in plant organic matter is currently used as a natural tracer of nitrogen acquisition efficiency. However, the δ¹⁵N value of whole leaf material does not properly reflect the way in which N is assimilated because isotope fractionations along metabolic reactions may cause substantial differences among leaf compounds. In other words, any change in metabolic composition or allocation pattern may cause undesirable variability in leaf δ¹⁵N. Here, we investigated the δ¹⁵N in different leaf fractions and individual metabolites from rapeseed (Brassica napus) leaves. We show that there were substantial differences in δ¹⁵N between nitrogenous compounds (up to 30‰) and the content in (¹⁵N enriched) nitrate had a clear influence on leaf δ¹⁵N. Using a simple steady‐state model of day metabolism, we suggest that the δ¹⁵N value in major amino acids was mostly explained by isotope fractionation associated with isotope effects on enzyme‐catalysed reactions in primary nitrogen metabolism. δ¹⁵N values were further influenced by light versus dark conditions and the probable occurrence of alternative biosynthetic pathways. We conclude that both biochemical pathways (that fractionate between isotopes) and nitrogen sources (used for amino acid production) should be considered when interpreting the δ¹⁵N value of leaf nitrogenous compounds.     

Foraging niche shift maintains breeding parameters of a colonial waterbird during range expansion

Relating the effects of foraging niche variation to reproductive dynamics is critical to understand species response to environmental change. We examined foraging niche variations of the slender‐billed gull (Chroicocephalus genei), a nomadic colonial waterbird species during its range expansion along the French Mediterranean coast over a 16‐year period (1998–2013). We investigated whether range expansion was associated with a change in chick diet, breeding success, and chicks body condition. We also examined whether breeding success and chicks body condition were explained by diet and colonial characteristics (number of pairs, laying phenology, habitat, and locality). Diet was characterized using dual‐stable isotopic proxies (δ¹³C and δ¹⁵N) of feather keratin from 331 individuals subsampled from a total of 4,154 chicks ringed and measured at 18 different colonies. δ¹³C decreased and δ¹⁵N increased significantly during range expansion suggesting that chicks were fed from preys of increasing trophic level found in the less salty habitat colonized by the end of the study period. Niche shift occurred without significant change of niche width which did not vary among periods, habitats, or localities either. Breeding success and chick body condition showed no consistent trends over years. Breeding success tended to increase with decreasing δ¹³C at the colony level while there was no relationship between stable isotope signatures and chick body condition. Overall, our results suggest that even if range expansion is associated with foraging niche shift toward the colonization of less salty and more brackish habitats, the shift had marginal effect on the breeding parameters of the Slender‐billed gull. Niche width appears as an asset of this species, which likely explains its ability to rapidly colonize new locations.     

Amino acid δ15N underestimation of cetacean trophic positions highlights limited understanding of isotopic fractionation in higher marine consumers

Compound‐specific stable isotope analysis (CSIA) of amino acids (AAs) has been rapidly incorporated in ecological studies to resolve consumer trophic position (TP). Differential ¹⁵N fractionation of “trophic” AAs, which undergo trophic ¹⁵N enrichment, and “source” AAs, which undergo minimal trophic ¹⁵N enrichment and serve as a proxy for primary producer δ¹⁵N values, allows for internal calibration of TP. Recent studies, however, have shown the difference between source and trophic AA δ¹⁵N values in higher marine consumers is less than predicted from empirical studies of invertebrates and fish. To evaluate CSIA‐AA for estimating TP of cetaceans, we compared source and trophic AA δ¹⁵N values of multiple tissues (skin, baleen, and dentine collagen) from five species representing a range of TPs: bowhead whales, beluga whales, short‐beaked common dolphins, sperm whales, and fish‐eating (FE) and marine mammal‐eating (MME) killer whale ecotypes. TP estimates (TP_CSIA) using several empirically derived equations and trophic discrimination factors (TDFs) were 1–2.5 trophic steps lower than stomach content‐derived estimates (TP_SC) for all species. Although TP_CSIA estimates using dual TDF equations were in better agreement with TP_SC estimates, our data do not support the application of universal or currently available dual TDFs to estimate cetacean TPs. Discrepancies were not simply due to inaccurate TDFs, however, because the difference between consumer glutamic acid/glutamine (Glx) and phenylalanine (Phe) δ¹⁵N values (δ¹⁵N_Glx‐Phe) did not follow expected TP order. In contrast to pioneering studies on invertebrates and fish, our data suggest trophic ¹⁵N enrichment of Phe is not negligible and should be examined among the potential mechanisms driving “compressed” and variable δ¹⁵N_Glx‐Phe values at high TPs. We emphasize the need for controlled diet studies to understand mechanisms driving AA‐specific isotopic fractionation before widespread application of CSIA‐AA in ecological studies of cetaceans and other marine consumers.